A Revision of Aganosma (Blume) G. Don (Apocynaceae) Author(s): David J. Middleton Source: Kew Bulletin, Vol. 51, No. 3 (1996), pp. 455-482 Published by: Springer on behalf of Royal Botanic Gardens, Kew Stable URL: http://www.jstor.org/stable/4117024 . Accessed: 16/07/2011 02:46 Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at . http://www.jstor.org/action/showPublisher?publisherCode=kew. . Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected].

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A revision of Aganosma (Blume) G. Don (Apocynaceae) DAVID

J. MIDDLETON'

Summary.The Asian genus Aganosmais revised. Eight species are recognised of which one has two varieties.The complex nomenclaturalissuesin the genus are discussed.

INTRODUCTION

Aganosmawas first described by Blume (1826) as a section of Echitesand raised to generic status by G. Don (1837) with a number of species based on specimens in Wallich's herbarium. Unfortunately considerable nomenclatural problems arose in the genus very early in its history. De Candolle (1844) split the genus into two sections Amphineurionand Meiadenia. The former consisted of four species which have now all been included in synonymy of Aganosma marginata. Bentham

& Hooker

(1876)

included

Aganosma as two

sections within Ichnocarpus but this was not followed by Hooker (1882) who reinstated the genus. Pichon (1948, 1950) raised de Candolle's two sections to generic level as Amphineurion and Aganosma but this has not been generally accepted, based as it is on an artificial weighting placed on the shape and indumentum of the area where the stamens cohere to the pistil head (see Leeuwenberg 1988; Middleton 1994). Aganosma belongs in the subfamily Apocynoideae, often previously called Echitoideae,

of

the

family

Apocynaceae.

The

position

of

Aganosma

in

the

subclassification of the family has been treated by a number of authors (De Candolle 1844; Bentham & Hooker 1876; Hooker 1882; Pichon 1950; Tsiang & Li 1977; Ly 1986). These treatments vary considerably reflecting a general lack of knowledge and research into the subclassification of the whole subfamily. Until a more thorough and satisfactory system for the Apocynoideaeis proposed I would prefer to indicate only that Aganosma would appear to be most closely related to Ichnocarpus, Trachelospermum,Epigynum and Chonemorpha.

Ly (1986) suggested the type species of the genus is Aganosma caryophyllataG. Don. Blume's concept of Echites caryophyllatais the type species of his section Aganosmaand therefore of the genus. This is now to be treated as a synonym of A. wallichiiwhich should, therefore, be the type species of the genus.

Accepted for publication January 1996. Department of Botany, Trinity College, Dublin 2, Ireland. Present address: Royal Botanic Garden, Inverleith Row, Edinburgh, EH3 5LR, U.K

455

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KEW BULLETIN VOL. 51(3) MATERIALS AND METHODS

Herbarium material was studied from the following herbaria: A, AAU, ABD, B, BK, BKF,BM, BO, BR, BRI, C, CGE, DBN, E, G, G - DC, GH, H, IBSC, K, KEP,KLU, K-W, KYO, L, LAE, M, MEL, MO, P, PE, S, SING, SINU, TCD, TI, U, UPS, US, W, WRSL, Z (Holmgren et al. 1990). All specimens cited have been seen unless otherwise stated. The dimensions given in the descriptions are for dried material except for the gynoecium and androecium characters which are for flowers rehydrated with water. Any type specimens associated with taxa described by Tsiang previously cited for LU or SYS now appear to be housed in IBSC and the holotypes are, therefore, cited IBSC. SYSTEMATIC TREATMENT

Aganosma (Blume) G. Don, Gen. Syst. 4: 77 (1837). Echitessect. AganosmaBlume, Bijdr. 1040 (1826). Aganosmasect. MeiadeniaA. DC., Prod. 8: 432 (1844). Ichnocarpussect. Meiadenia (A. DC.) Bentham & Hooker, Gen. P1. 2: 717 (1876). Type species: Aganosma wallichii G. Don.

Aganosmasect. AmphineurionA. DC., Prod. 8: 433 (1844). Ichnocarpussect. Amphineurion(A. DC.) Bentham & Hooker, Gen. Pl. 2: 717 (1876). Aganosmasect. AmphyneurionBoerl., Handl. Fl. Ned. Ind. 2: 399 (1899) sphalm. Amphineurion(A. DC.) Pichon, Bull. Soc. Bot. France 95: 215 (1948). Type species: Aganosma marginata (Roxb.) G. Don.

Climbers or scramblers; producing latex. Stems often lenticellate, sometimes densely so, glabrous or pubescent. Leavesopposite, those of a pair equal, petiolate, coriaceous to papery, entire. Inflorescencea terminal panicle or a pseudoaxillary panicle. Flowers5-merous, actinomorphic, usually fragrant. Sepallobes free; colleters present or absent, only on inside corners or in a row inside. Corollalobes in bud overlapping to the right; tube narrowly cylindrical or somewhat inflated below, widening slightly at the point of stamen insertion into the upper tube, then with spreading lobes; lobes elliptic or obovate-falcate. Stamensincluded in the corolla tube, attached in a ring to the pistil head; anthers subsessile, narrowly triangular, apex acuminate, base sagittate, sterile at apex and base. Disk usually of 5 fused, occasionally free, lobes, narrower at top and surrounding ovary or of 5 rounded lobes shorter than ovary. Ovary of 2 separate carpels united into a common style, superior, ovoid, glabrous or pubescent; ovules numerous; style glabrous or, rarely, pubescent; pistil head ovoid with a short sharp projection on top. Fruit of paired follicles, divergent or not, linear and narrow or somewhat fusiform, longitudinally dehiscent. Seedsnarrow elliptic, flattened, glabrous, with an apical unbeaked coma. Eight species from India to China and southwards to the Philippines and western Indonesia.

A REVISION OF AGANOSMA (APOCYNACEAE)

457

KEY TO SPECIES OF Aganosma

1. Leaves with a strong intramarginal nerve; sepals shorter than to as long as the corolla tube; sepals with continuous row of colleters; disk shorter than to as 5. A. marginata long as ovary ................................... Leaves without a strong intramarginal nerve; sepals longer than the corolla tube; sepals with colleters only in the corners or absent; disk longer than ovary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . 2 2. Corolla throat glabrous ....................................... 3 Corolla throat pubescent ...................................... 5 3. Leaves with 2 - 4(- 6) pairs of lateral nerves, very strongly ascending at an angle of 25 - 55'; stamens inserted at 0.4 - 0.5 of tube length .... 3. A. dichotoma Leaves with 5 - 13 pairs of lateral nerves, not strongly ascending, 55 - 75'; stamens inserted at 0.2- 0.4 of tube length ....................... 4 4. Corolla lobes elliptic, acuminate; tube 4.1 - 8 mm long ................. ...... 2. A. cymosa var. cymosa ........................ ..... Corolla lobes obovate, flat topped, possibly acute in one corner; tube 8.3 - 14 mm long .................................. 8. A. wallichii ..... 5. Corolla lobes obovate, apex rounded or with one acute corner ............ A. schlechteriana ........................................6. 6 Corolla lobes elliptic, apex acuminate or acute ....................... 6. Corolla lobes 2.8- 8.2 mm long ................................. 7 Corolla lobes 11 -41 mm long .................................. 8 7. Inflorescence (3.1 -)5.3 - 8 cm long; China, Burma, Thailand - - 1. A. breviloba Inflorescence 1.1 - 4.5 cm long; southern India, Sri Lanka ............... A. cymosa var. elegans ....................................2. 8. Ovary glabrous .......................................4. A. gracilis ................... A. siamensis Ovary pubescent ..................7. 1. Aganosma breviloba Kerr, Bull. Misc. Inform., Kew 1937: 92 (1937); Kerr, Fl. Siam. En. 2: 468 (1939). Type: Thailand, Chiang Mai, Doi Sutep, Kerr 3211 (lectotype K, designated here; isotypes ABD, BM, E, K, L, P, TCD). Aganosma lacei Raizada, Indian Forester 68: 363 (1942). Type: Lace 5251 (holotype DD n.v.; isotypes E, K). Stems densely to sparsely puberulent when young, generally becoming glabrous with age. Leaves:petiole 6 - 12 mm long; blade elliptic, apex acuminate, base cuneate to obtuse, 5.5 - 13.5 x 2.2 - 6.4 cm, 2 - 6 x as long as wide, 5 - 8 pairs of lateral nerves, glabrous or with few strigose hairs on abaxial midrib, petiole and margin at base. Inflorescencea delicate terminal panicle, pubescent, 3.1 - 8 cm long; pedicels 3 - 10 mm long. Sepalslinear, 10 - 12 x 1.5 - 3.7 mm, 3.4 - 6.7 x as long as wide, longer than corolla tube, pubescent, with narrow colleters in the corners. Corollawhite; tube 6.7 10 mm long; lobes narrow elliptic, acuminate, 5 - 8.2 x 1.8 - 2.6 mm, 2.4- 3.4 x as long as wide, 0.6 - 0.9 x as long as tube; short pubescent outside, longer pubescent in throat and in 5 rows inside. Stamensinserted in tube 2- 3.8 mm from base, 0.2 - 0.4 of tube length; anthers 4.4- 6 x 0.9 - 1.1 mm, 4.9- 6 x as long as wide. Disk narrow

458

VOL. 51(3) KEWBULLETIN

at top, 5-dentate, 1.1 - 2.2 mm long, 1.6 x as long as ovary. Ovary0.9 - 1.4 mm long, pubescent to very sparsely pubescent; style 2 - 2.4 mm long; pistil head 2.7 - 3.8 mm long. Fruit unknown. Fig. 1. Map 1. DISTRIBUTION. China, Burma, Thailand. HABITAT. In evergreen forest, 750 - 1220 m. CHINA. Guizhou, Cavalerie & Fortunat 2156 (P) BURMA. Shan State, King's Collector 651 (K); Mandalay, Maymyo, Lace 5251 (E,

K-

type of A. lacei). Chiang Mai, Doi Sutep, Kerr 3211 (ABD, BM, E, K, L, P, TCD -

THAILAND.

type

of A. breviloba).

2. Aganosma cymosa (Roxb.) G. Don, Gen. Syst. 4: 77 (1837); A. DC., Prod. 8: 433 (1844); Hook. f., Fl. Brit. Ind. 3: 665 (1882); Watt, Products of India 1: 129 (1889) pro parte; Boerl., Handl. Fl. Ned. Ind. 2: 399 (1899); Brandis, Indian Trees 464 (1906); Tsiang, Sunyatsenia 4 (1939) 35. EchitescymosaRoxb., Fl. Ind. 2: 16 (1832); Wight, Ic. P1. 2: t. 395 (1843); Voigt, Hort. Sub. Calc. 523 (1845). Type: Bangladesh, Sylhet, Wallich1654a proparte (lectotype K -W [as 1654.1], designated here; isotypes BM, BR, E, G, K, L, M, MEL, TCD, W). KEY TO VARIETIES OF A. cymosa

1. Corolla glabrous in throat ............................ Corolla densely pubescent in throat .....................

a. var. cymosa b. var. elegans

a. var. cymosa Aganosma cymosavar. glabraA. DC., Prod. 8: 433 (1844). Type: Bangladesh, Sylhet, Wallich1654a pro parte (lectotype G - DC, designated here; isotypes BM, BR, E, G, K, K - W, L, S, W).

Aganosmacymosavar. lanceolataHook. f., Fl. Brit. Ind. 3: 665 (1882). Type: Cultivated in Calcutta, Wallich1654B (lectotype K - W [as 1654.2], designated here; isotypes K, L, W). AganosmaharmandianaPierre ex Spire, Contr. Apocyn. 108 (1905); Pitard, Fl. Gen. Indo-Chine 3: 1225 (1933); Tsiang, Sunyatsenia 4: 38 (1939); Tsiang, Bull. Fan Mem. Inst. Biol. 9: 24 (1939); Kerr, Fl. Siam. En. 2: 468 (1939); Tsiang & P. T. Li, Fl. Rei. Pop. Sin. 63: 183 (1977); J. Ying & P. T. Li, Fl. Yunnanica 3: 536 (1983); Ly, Feddes Repert. 97: 658 (1986). Type: Laos, Lakhone, Harmand 5241 (lectotype P, designated here; isotypes P, TCD). Aganosma cymosavar. fulva Craib, Feddes Repert. Nov. Spec. Regni Veg. 12: 393 (1913). Type: Burma, Tavoy, Bowachoung, Meebold 15365 (lectotype K, designated here; isotype K). Stems densely tomentose or strigose when young, becoming glabrous with age. Leaves:petiole 0.5 - 2.8 cm long; blade elliptic to weakly obovate, apex acuminate to obtuse, base cuneate to rounded, 4- 16 x 1.9 - 9.6 cm, 0.7 - 3.9 x as long as wide, 5 - 10

A REVISION OF AGANOSMA (APOCYNACEAE)

FIG. 1. Aganosma breviloba.A Habit; B Flower; C Dissected flower. Drawn from Kerr3211 by Holly Nixon.

459

460

KEWBULLETINVOL.51(3)

.,........ ... .

.,,7

Z.

i/

I

J•

,. ..l

,...

.o;

?

AL0 Alt

1

;I

44j 44\

MAP 1. Distribution elegans (A) .

of Aganosma breviloba (u);

?~~

Aganosma cymosa var. cymosa (0) and Aganosma cymosa var.

pairs of lateral nerves, usually densely tomentose abaxially and often adaxially, sometimes becoming glabrous, rarely only sparsely pubescent. Inflorescence congested, terminal, tomentose, 2.4 - 9.2 cm long; pedicels 2.3 - 6.5 mm long. Sepals narrow ovate to lanceolate, 7.1 - 16 x 1.2 - 3.1 mm, 2.7 - 7.6 x as long as wide, longer than corolla tube, tomentose, with few colleters in corners or absent. Corolla white; tube 4.1 - 8 mm long; lobes elliptic, acuminate, 4.7 - 13 x 1.4 - 4.1 mm, 1.9 4.3 x as long as wide, 0.8 - 2.5 x as long as tube; sparsely to densely tomentose outside, glabrous in throat, pubescent in rows behind anthers. Stamens inserted 1 - 2.7 mm from base of corolla tube, 0.2 - 0.4 of corolla tube length; anthers 3.6 - 4.9 x 0.7 - 1 mm, 4.6 - 6.7 x as long as wide. Disk narrower at top, 5-dentate, 1 - 1.9 mm long, 1.4 - 3.7 x as long as ovary. Ovary 0.3 - 0.8 mm long, glabrous or pubescent; style 0.7 - 1.7 mm long; pistil head 1.6 - 3 mm long. Fruit fusiform or linear terete, often curved, densely hirsute, 8.1 - 37 x 6 - 9.3 mm. Seeds 12.3 - 22.5 x 3.7 - 6.8 mm; coma 2.5 - 5.1 cm long. Fig. 2. Map 1.

A REVISION OF AGANOSMA (APOCYNACEAE)

461

FIG. 2. Aganosma cymosa var. cymosa A Habit; B Flower; C Dissected flower; D Fruit; E Seed. Drawn by Holly Nixon; flowering portion from Maxwell 72-291, fruiting portion from Kerr 2778.

462

KEWBULLETIN VOL. 51(3)

DISTRIBUTION. India (Assam, Nagaland), Bangladesh, Burma, China, Thailand, Laos, Cambodia, Vietnam. HABITAT. In evergreen, mixed or deciduous forest or thicket, to 1300 m. Selection of the 40 specimens studied: INDIA. Cultivated in Calcutta, Wallich1654 B (K, K - W, L, W - type of A. cymosa var lanceolata); Meghalaya, Cherrapunjee, Khasi Hills, Chand 5581 (L); Nagaland, Emilorni, Naga Hills, Bor5063 (K). BANGLADESH. Sylhet, Wallich1654 A (BM, BR, E, G, K, L, MEL, TCD - type of Echitescymosa);Wallich1654.1 or a (BM, BR, E, G, K, L, S, W - type of A. cymosavar. glabra). BURMA. Myitkyina, Nsop Zup, Mikee6241 (K, P, US); Shan Hills, Terai, Collett825 (K - Syntype of A. cymosavar. fulva); Tavoy, Bowachoung, Meebold15365 (K lectotype of A. cymosavar.fulva); Eastern Tenasserim, Kerr21592 (BM, K); Bhamo district Gyi4521 (K). CHINA. Yunnan: Szemao, Henry 12761 (E, MO, K); Che-li Hsien, Wang 75559 (A); Dah-meng-lung, Che-li Hsien, Wang77601 (UPS), Wang78023 (A); Mong-hain, Che-li Hsien, Wang79878 (A); Mienning, Yewanshui, Yu 17686 (A). THAILAND.Chaiyaphum, Ban Nam Phrom, Geesinket al. 6840 (AAU, BKF, C, K, L); Chanthaburi, Doi Soi Dao Nue, Geesinket al. 6750 (AAU, BKF, C, K, L, P ); Chiang Mai, Doi Sutep, Kerr2778 (ABD, BM, E, K, TCD); Chon Buri, Si Racha, Marcan 186 (BM, K); Chon Buri, Sattahip, Tong Brong, Maxwell 71418 (AAU); Lampang, Me Yom, Winit 1828 (BKF,BM, K); Saraburi, Menam Sak, Kerr7060 (BM, E, K, L, P, TCD); Sukhothai, Maxwell72-291 (AAU). CAMBODIA. Phnom Penh, Bejaud55 (P, TCD). LAOS. Lakhone, Harmand 5241 (P, TCD - type of A. harmandiana);Phon Soun, Luang Prabang, Poilane 20097 (P); Tchepone between a Trup et a Pril, Poilane 13475 (P). VIETNAM. S.1l.Mueller24 (P); Baria, Poilane 672 (P). NOTE. The type number of Echites cymosa, Wallich 1654.1 in the Wallich herbarium, closely fits the description given by Roxburgh (1832). This plant has densely tomentose leaves and a glabrous corolla throat. De Candolle's var. glabra, also typified by Wallich1654, is represented by some glabrous forms of this species under this number. This plant also has a glabrous corolla throat but slightly larger flowers than the tomentose form. The glabrous form looks superficially more like Wallich1655 and 1656, respectively labelled as Echitesconfertaand Echiteselegans.The tomentose form is very similar to a number of taxa described from further east, particularly A. harmandiana.It has generally been accepted that Echitesconfertaand E. elegans and combinations based on them are synonyms of Aganosma cymosa. Unfortunately the much more common southern Indian and Sri Lankan variety seems to have been taken as the norm and the similarity between the types of Echites cymosa,Aganosmaharmandianaand A. cymosavar.fulva has been overlooked.

b. var. elegans (G. Don) Hook.f, Fl. Brit. Ind. 3: 665 (1882); Boerl., Handl. Fl. Ned. Ind. 2: 399 (1899); Tsiang, Sunyatsenia 4: 36 (1939); Huber in Dassanayake & Fosberg, Fl. Ceylon 4: 55 (1983).

A REVISION OF AGANOSMA (APOCYNACEAE)

463

Aganosma elegans G. Don, Gen. Syst. 4: 77 (1837); A. DC., Prod. 8: 433 (1844); Wight, Ic. P1. 4: t. 1304 (1850); Thwaites, Enum. P1. Zeyl. 194 (1860). EchiteselegansWall., Numer. List 1656 (1829) nom. nud. Type: India, s.l., Wallich1656 (lectotype K - W [as 1656.1], designated here; isotypes BM, E, G, K, L, SING). Aganosma cymosa(Roxb.) G. Don: Watt, Products of India 1: 129 (1889) pro parte; Trimen, Handbook Fl. Ceylon 3: 139 (1895); Cooke, Fl. Pres. Bombay 2: 140 (1908); Gamble, Fl. Pres. Madras 5: 819 (1923); Huber in Dassanayake & Fosberg, Fl. Ceylon 4: 55 (1983). Aganosmadoniana Wight, Ic. P1. 4: t. 1306 (1848); Dalzell & Gibson, Bombay Fl. 146 (1861). Type: Wight115 (holotype K). Aganosma cymosavar. conferta(G. Don) Hook. f., Fl. Brit. Ind. 3: 665 (1882). Echites confertaWall., Numer. List 1655 (1829) nom. nud. AganosmaconfertaG. Don in Loud., Encycl. P1.Suppl. 2: 1312 (1855) nom. nud. Type: India, s.l., Wallich1665 (lectotype K - W, designated here; isotypes BM, G, K). ToxocarpuseriocarpusHook. f., Fl. Brit. Ind. 4: 15 (1885). Type: India, Courtallan, Wight2234 (holotype K). EchitesbracteataWall., Num. List 1656 (1829) proparte nom. nud. [1656.B in K - W] Stems glabrous, sparsely pubescent, strigose or tomentose. Leaves:petiole 0.6 - 2 cm long; blade elliptic or, rarely, obovate, apex acuminate to acute, base cuneate to rounded,

2.3 - 11.7 x 0.8 - 6.7 cm, 1.4 - 4.4 x as long as wide, 4 - 8 pairs of lateral

nerves, often strongly ascending, glabrous or pubescent on midrib and petiole. Inflorescence congested, terminal, tomentose, 1.1 - 4.5 cm long; pedicels 2.4 - 10 mm long. Sepalsnarrow lanceolate, 4.8 - 11 x 0.6 - 2.6 mm, 3.5 - 10.2 x as long as wide, longer than corolla tube, tomentose, few colleters in corners or absent. Corolla white; tube 4.3 - 8 mm long; lobes elliptic, acuminate,

2.8 - 6.5 x 1.2 - 3 mm, 1.8 -

3.1 x as long as wide, 0.6 - 1.3 x as long as tube; tomentose outside, densely pubescent in throat and in 5 rows in tube inside. Stamensinserted at 1.5 - 2.4 mm from base of corolla tube, 0.3 - 0.4 of tube length; anthers 2.5 - 4.3 x 0.5 - 0.8 mm,

3.1 - 6.4 x as long as wide. Disk narrower at top, 5-dentate or, rarely, with some free

lobes, 0.9 - 1.7 mm long,

1.5 - 3.3 x as long as ovary. Ovary 0.4 - 0.6 mm long,

densely pubescent; style 1 - 1.8 mm long; pistil head 2 - 2.8 mm long. Fruit fusiform or terete, tomentose to puberulent, 9 - 30 x 0.55 - 0.8 cm. Seeds17 - 30.2 x 4.3 - 5.5 mm; coma 3.1 - 7 cm long. Map 1. DISTRIBUTION. Southern India, Sri Lanka.

HABITAT.In evergreen or deciduous forest or scrub to 1550 m. Selection of the c. 70 specimens studied: INDIA. s.l. Wallich1655 (BM, G, K - type of Echites cymosavar. conferta); Wallich 1656 (BM, E, G, K, L, SING - type of Aganosmaelegans). Karnataka:Attur District, Salem, Periakalrayans, Chinnakkenattharu Konavari Matthew RHT 15255 (K); Hassan District, stream between Devalkere & Devarunde, RamamoorthyHFP 1600 (BO, US). Kerala: environs of Munnar, Blasco 8009 (L); Courtalan, Wight531 (E). Maharashtra: s.l. Gibsons.n. (L, M), Youngs.n. (BM). Tamil Nadu: Pulneyo, Bourne 1599 (K); Cuddapah District, Gamble 20979 (K); Kaguchi, Lawson 12874 (K); Kodaikanal Region, Palni Hills, Anglade s.n. (A); Billigirangan, Barnes 321 (A, US);

464

KEWBULLETIN VOL. 51(3)

Nilgirri Hills, Cleghorns.n. (E); Palni Hills, Sauliere 800 (K); Tirunelveli Dept., Nadukani Road, Shetty 27973 (L); Courtallan,

Wight 2234 (K -

type of Toxocarpus

eriocarpus).Kerala: Environs of Munnar, Blasco8009 (L). SRI LANKA. Central: Matale District, 9.5 miles ENE of Dambulla,

Davidse &

Sumithraarachchi 8135 (K, KLU, L, MO). Eastern: Trincomalee District, TrinHabarana road, Sohmer8178 (US); Batticaloa District, Gunners Quoin, Tirvengadum et al. 234 (K, L, P, US). North Central: Anuradhapura District, Between Magoduwewa and Huluwewa, Balakrishnan NBK 1063 (K, MO, P, US); Anuradhapura District, Ritigala Strict Natural Reserve, Jayasuriya& Premadasa1633 (L, US). Uva: Monaragala District, Northern slope of Kataaragama Peak, Huber65 (K, MO, US). North Western: Kurunegala, Kostermans24937 (G, Z). 3. Aganosma dichotoma

(Roth in Roem. & Schult.) K. Schum. in Engler & Prantl, Nat.

Pflanzenfam. 4(2): 173 (1895); Gamble, Fl. Pres. Madras 5: 819 (1923). Tsiang, Sunyatsenia 4: 40 (1939); Mooney, Suppl. Bot. Bihar & Orissa 225 (1950); Maheshwari & Singh, Dict. Econ. Pl. India 5 (1964). Echites dichotomaRoth in Roem. & Schult., Syst. Veg. 4: 393 (1819) non Thunberg (1819) nec Humb., Bonpl. & Kunth (1819); Roth, Nov. Pl. Sp. 136 (1821). EchitesheyneiSprengel, Syst. Veg. 632 (1824). dichotomaG. Don, Gen. Syst. 4: 76 (1837). Type: Wallich1651 (neotype Chonemorpha K-W [as 1651.1]) Aganosma caryophyllataauct. non G. Don: Graham, Cat. Bomb. Pl. 113 (1839); A.DC., Prod. 8: 432 (1844) pro parte;Voigt, Hort. Sub. Calc. 523 (1845); Dalzell & Gibson, Bombay Fl. 51 (1861); Hook. f., Fl. Brit. Ind. 3: 664 (1882); Watt, Products of India 1: 129 (1889); Cooke, Fl. Pres. Bombay 2: 140 (1904); Brandis, Indian Trees 464 (1906); Haines, Fl. Chota Nagphur 428 (1919); Haines, Bot. Bihar & Orissa 4: 545 (1922); Tsiang, Sunyatsenia 2: 154 (1934); Kanjilal, Fl. Assam 3: 268 (1939). AganosmaroxburghiiG. Don, Gen. Syst. 4: 77 (1837) excluding type; Wight, Ic. Pl. 2: t. 440 (1843).

Echites caryophyllataauct. non Roxb.: Sims, Bot. Mag. 44: t. 1919 (1817); Roem. & Schult., Syst. Veg. 4: 392 (1819); Roth, Nov. Pl. Sp. 133 (1821). Aganosmablumeiauct. non A. DC.: Wight, Ic. P1. 4: t. 1305 (1850). Stems glabrous, sparsely strigose or tomentose. Leaves:petiole 5 - 28 mm long; blade ovate to elliptic, apex acuminate, rarely to obtuse, base rounded, rarely obtuse or weakly cordate, 2.8 - 14.2 x 1.3 - 4.8 cm, 1.3 - 2.8 x as long as wide, 2 - 4(- 6)

pairs of lateral nerves, very strongly ascending, glabrous, pubescent only on abaxial midrib or pubescent all over abaxially. Inflorescence lax, terminal, sparsely to densely 3 cm 20.5 pubescent, long; pedicels 2.8 23 mm long. Sepalsnarrow lanceolate, 12 - 25 x 1 - 3.7 mm, 4 - 15 x as long as wide, longer than corolla tube, tomentose,

with colleters in the corners inside. Corolla white; tube 8 - 13 mm long; lobes obovate, falcate, 7.3- 20 x 4.6- 14 mm, 1 - 2.4 x as long as wide, 0.8- 1.7 x as long as tube; very sparsely puberulent to puberulent outside, glabrous in throat, pubescent in 5 rows behind anthers. Stamensinserted at 3.5 - 6.2 mm from base of corolla tube, 0.4 - 0.5 of tube length; anthers 4.4 - 5.8 x 0.9 - 1 mm, 4.7 - 6.1 x as

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long as wide. Disk narrower at top, 5-dentate or with some, but not all, lobes free, 0.9 - 1.9 mm long, 1.3 - 2.4 x as long as ovary. Ovary 0.7 - 0.8 mm long, pubescent; style

2.8 - 6.4 mm long; pistil head 1.9 - 2.7 mm long. Fruit fusiform, divergent to parallel, tomentose

to glabrous, 6.3 - 21 x 0.7 - 1 cm. Seeds 10.5 - 18 x 4.3 - 6.6 mm;

coma 1.4 - 3 cm long. Fig. 3. DISTRIBUTION. Hooker (1882) noted that it was difficult to give a natural distribution for this species as it is widely cultivated all over India. I have seen only two collections, Haines 4877 and Haines 4880, which would seem to have been collected in the wild. These were both collected near Puri in Orissa in India.

Selection of the c. 50 specimens studied (all cultivated): INDIA. Andra Pradesh: Cuddapah, Beddome 5145 (B, BM). Bihar: Munger, Hamilton 732 (E). Calcutta Botanic Garden: Pierre3576 (A, P); ibid., Pierres.n. (P, TCD). Chota Nagpur: Hazaribagh District, F H. W Kerrs.n. (BM). Tamil Nadu: Cuddapah District, Chengelet Pass, Gamble21222 (K); Godavari District, Rampa Hill, Ramaswami 1535 (K). Nicobar Islands: Galathea, Wallich 1035 (C). Orissa: Cuddapee Hills, Beddome5140 (BM); Cuttack, Beddome5141 (BM); Angul, Mooney 500 (K). Tamil Nadu: Kondapalli, Gamble 18583 (K). Uttar Pradesh: Benares, Ayurveda Botanic Garden, Schwabes.n. (B). Maharashtra: Bombay, Dalzells.n. (K). INDONESIA. Java: s.l., Zollinger 3154 (BM, G, P, S). MALAYSIA.Penang: Unknown s.n. (E). SINGAPORE. Botanic Gardens, Jameson 699 (E).

EchitesdichotomaRoth in Roem. & Schult. is the basionym of Aganosma dichotoma (Roth) K. Schum. (in Engler & Prantl, Nat. Pflanzenfam. 4(2): 173. 1895), a frequently cultivated species of Aganosma from India. There has, however, been confusion over the correct name for this species with the alternative name Aganosma caryophyllataG. Don also having been used. However, Aganosma caryophyllata,as originally described and typified by Don, contains no element which belongs to the taxon in question. Roxburgh published Echitescaryophyllatain 1814 (Hort. Beng. 19. 1814) and cited Rheede's Hort. Mal. 9: t. 14 under it. Roxburgh did not include a description, but the name is validated by reference to the thorough description and illustration in Rheede (1689). The plant illustrated and described by Rheede, however, turns out to be a Kamettiaand Echitescaryophyllatahas been used as the basionym for Kamettia caryophyllata(Roxb.) Nicolson & Suresh (in Taxon 35: 354. 1986). In 1817 Sims (Bot. Mag. 44: 1919. 1817) gave a description of E. caryophyllatabut used the Rheede name of "Tsjeria-pu-pal-valli"and the citation of Hort. Mal. 7: p.103. t. 55, a plant which, from the description and the illustration, is quite clearly an Aganosma. He cited Roxburgh as ined. and noted that Brown (Asclep. 2. 1810) suggested that this plant (i.e. the Rheede 7: t. 55 plant) might deserve generic status outside of Echites. Blume (Bijdr. 1041. 1826) placed E. caryophyllataRoxb. in his new section Aganosma.Although he attributed the Indonesian plant he described to Roxburgh it is very clear from his description that he is describing an Aganosma and not a Kamettia.He also cited the illustration in Rheede's Hort. Mal. 7: t. 55.

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FIG. 3. Aganosma dichotoma A Habit; B Flower; C Flower bud; D Dissected flower. Drawn from Mooney 500 by Holly Nixon.

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In 1832 Roxburgh described Echites caryophyllatain his Flora Indica. The plant described is very clearly an Aganosma but he still cites the Rheede name for the Kamettia,"Kemettivalli", albeit with the wrong Hort. Mal. citation of 9: t. 135 instead of 9: t. 14. This is clearly a printing error as 135 does not exist in the work. G. Don (Gen. Syst 4: 77. 1837) transferred both E. caryophyllataRoxb. and E. caryophyllatasensu Blume to Aganosma. He believed the Blume name had priority over the Roxburgh name, presumably because it was the Blume plant which was the basis for section Aganosma and, therefore, for the genus Aganosma.Echites,as now delimited, is confined to the Americas and Aganosmais one of a number of generic segregates. Don transferred Echites caryophyllatasensu Blume to Aganosma as A. caryophyllataand E. caryophyllataRoxb. to the new name of A. roxburghii.He cited Rheede, Hort. Mal. 7: 103, t. 55 under A. caryophyllataand Rheede, Hort. Mal. 9: t. 135, the incorrect Roxburgh citation of 9: t. 14, under A. roxburghii. He also included Sims' Echites caryophyllataunder A. roxburghii.Don's A. caryophyllatamust be taken as a new name and not a new combination as Blume's Echitescaryophyllata is not a valid basionym. It contains two distinct elements: the description of an Indonesian plant and the citation of the Rheede figure, Hort. Mal. 7: t. 55. A. de Candolle (Prod. 8: 432. 1844) decided that the Rheede element 9: t. 14 belonged with the epithet caryophyllataand also included A. roxburghiiin synonymy. He specifically excluded the Blume element for which he erected the new name Aganosma blumei,including Rheede, Hort. Mal. 7: t. 55, indicating that it grew in both Java and India. Wight (Ic. t. 1305. 1848) also described and illustrated A. blumeibut his citation was specifically excluded from A. blumeiby Miquel (Fl. Ind. Bat. 446. 1857). This plant most closely resembles A. cymosa. Hooker (Fl. Brit. Ind 3: 664. 1882) decided the problem species was to be called Aganosmacaryophyllata,which is the plant clearly described under Don's A. roxburghii and the plant illustrated as E. caryophyllataby Sims. Hooker noted the fact that there was considerable nomenclatural difficulty with the name he decided to use but believed that it should be kept because it was commonly used for this species particularly in herbaria and in botanic gardens. He excluded Roxburgh's 1832 Echites caryophyllata(which he said described A. calycina [= A. wallichii]), Rheede's, Hort. Mal. 7: t. 55 and Rheede's 9: t. 14 from this species and indicated no distribution outside of India. By doing so he excluded the types of both Echites caryophyllataRoxb. and AganosmacaryophyllataG. Don. Continued use of the epithet 'caryophyllata' by application for conservation with a new type would perpetuate the use of the confusing name. Don's Aganosma caryophyllatais to be typified by one of the two elements it contains: a drawing of Rheede's, 7: t. 55, which clearly resembles A. cymosa(Roxb.) G. Don, or by Blume's description of an Indonesian plant. Based as it is on Blume's concept of A. caryophyllata,the latter is more justifiable. Blume's Echitescaryophyllata is more than likely Aganosmawallichii, the only Aganosmain Indonesia which would fit the description. A. blumei A. DC., the new name for Blume's plant, is still, however, typified by Rheede 7: t. 55, the Indian plant resembling A. cymosa,and the unknown Indonesian plant, both of which have already been used to typify Don's A. caryophyllataand is, therefore, an illegitimate name. Don's Aganosmacaryophyllatais

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clearly not the one of Hooker and as generally understood, this being Don's description,

but not typification, of A. roxburghii. A. caryophyllata G. Don should be

synonymised under A. wallichii or A. cymosa,preferably the former as the author clearly had the Indonesian element in mind in the description. K. Schumann (in Engler & Prantl, Nat. Pflanzenfam. 4(2): 173. 1895) published the combination Aganosma dichotoma (Roth) K. Schum with A. caryophyllata G. Don in synonymy. This

was based on the belief that Echites dichotomaRoth (Nov. Pl. Sp. 136. 1821) is an earlier basionym than E. caryophyllata. Roth's species, however, was actually published by Roemer & Schultes in 1819, after Roth made his manuscript available to them, which leads to confusion of priority with regard to the homonyms E. dichotomaThunb. (1819) and E. dichotomaKunth (in Humb., Bonpl. & Kunth, Nov. Gen. 3: 217. 1819). The dates given in TL-2 for these three homonyms indicate that the Kunth name is later but that the relative dates of the other two are open to debate. The Thunberg name is given as 21st April and the Roemer & Schultes name as March -June. E. heyneiSpreng. (Syst. Veg. 1: 632. 1824) is a new name for E. dichotomaRoth in Roem. & Schult. and is cited in synonymy of A. caryophyllataby Hooker (1882). There is one Heynecollection listed in the Wallich Catalogue under Echitescaryophyllatabut there is no indication of whether this is indeed the type of E. dichotomaRoth in Roem. & Schult. or E. heyneiSpreng. This specimen does not seem to be in the Wallich herbarium and the original specimen, upon which Roth based the name, would have been deposited in Berlin. This specimen cannot now be traced and was presumably destroyed. It is likely, though, that Schumann did examine this specimen before it was destroyed and came to the conclusion that it was synonymous

with Aganosma caryophyllata as generally

used. Echites dichotoma

Thunb. is not a currently used name and, therefore, it seems sensible to retain the Roth version until such a time as more definite dates are available and it is ascertained whether conservation is needed or not. The main argument against the use of Echitesdichotomais that the name Aganosma caryophyllatais a more widely accepted name for this taxon. Although this is indisputable,

Aganosma dichotoma, based on Echites dichotoma, has been used by

Gamble (Fl. Madras 819. 1923) and many herbarium specimens bear his identification.

Aganosma dichotoma is preferable to A. caryophyllata as it avoids the use

of the extremely confusing name and avoids the need to reject Echites dichotoma Roth and Echites heyneiSpreng., the latter an indisputably older name even if E. dichotomaRoth is a later homonym. All the names associated with A. caryophyllata can be accommodated in synonymy of other species of Apocynaceae. 4. Aganosma gracilis Hook. f., Fl. Brit. Ind. 3: 664 (1882); Brandis, Indian Trees 464 (1906); Tsiang, Sunyatsenia 4: 37 (1939) pro parte. Type: India, Sikkim, Hookers.n. (lectotype K, designated here; isotypes K, P, GH [photo in E]). Stems sparsely lenticellate; glabrous or sparsely strigose. or obovate, apex acuminate, base cuneate to obtuse; petiole 4.9 - 11.7 x 2.2 - 5 cm, 1.8 - 3.4 x as long as wide; with 6- 9 glabrous or, more rarely, sparsely pubescent on petiole Inflorescence lax and delicate;

pubescent;

Leaves elliptic, oblong 7 - 16 mm long; blade pairs of lateral nerves, and abaxial midrib.

3.2 - 17 cm long; pedicels

0.7 - 3.6 cm

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long. Sepals narrow lanceolate;

469

1.3 - 2.8 cm long, 1.3 - 4 mm wide, 5.7 - 13.3 x as

long as wide, longer than corolla tube; pubescent; with colleters in the corners. Corollawhite; tube 6 - 22 mm long; lobes narrow elliptic or lanceolate, acuminate, 19 - 41 mm long, 3.6 - 8.2 mm wide, 4.5 - 7.1 x as long as wide; lobes 1 - 2.3 x as

long as tube; short pubescent outside, pubescent in throat and in five rows inside. Stamens inserted at 2 - 2.9 mm from base, 0.1 - 0.2 of tube length; anthers 5.4 - 6

mm long, 0.9 - 1.1 mm wide, 5.2 - 6.7 x as long as wide. Disk narrower at top, 5dentate;

1.3 - 2.1 mm long; 1.3 - 3.5 x as long as ovary. Ovary 0.6 - 1.5 mm long;

glabrous; style 1.2 - 1.6 mm long; pistil head 2.5 - 3.5 mm long. Fruit unknown. Fig. 4. Map 2. DISTRIBUTION.

Bhutan, India.

HABITAT. In forest, 600 - 1200 m.

BHUTAN. Mongar District, Saleng, BowesLyon 15114 (BM); Mirichoma Timpu, Cooper 1077 (BM). BURMA. King Tung, MacGregor 720 (E). INDIA. Meghalaya: Cherrapunjee, Khasi Hills Chand 5715 (L); Koelz 30011 (L); Lobb s.n. - syntype of A. gracilis). Sikkim: s.l., Hooker s.n. (E (photo), GH, K, P -

type of A. gracilis); Chonbutee, Darjeeling, Clarke12086 (B, BM); Nampok, Pantling 46626 (A, K); Legship, Stainton5375 (BM). 5. Aganosma marginata (Roxb.) G. Don, Gen. Syst. 4: 77 (1837); Wight, Ic. P1. 2: t. 425 (1843); Kurz, Forest Fl. Burma 2: 186 (1877); Hook. f., Fl. Brit. Ind. 3: 663 (1882); Watt, Products of India 1: 129 (1889); Boerl., Handl. Fl. Ned. Ind. 2: 399 (1899); Spire, Contr. Apocyn. 110 (1905); Brandis, Indian Trees 464 (1906); King & Gamble, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74: 495 (1907); Ridl., Fl. Mal. Pen. 2: 365 (1923); Parkinson, For. Fl. Andaman Isl. 207 (1923); Pitard, Fl. Gen. IndoChine 3: 1222 (1933); Kerr, Fl. Siam. En. 2: 468 (1939); Backer & Bakh. f., Fl. Java 2: 237 (1965); Gangopadhyay & Chakraborty,J. Econ. Tax. Bot. 16 (1992) 29. EchitesmarginataRoxb., Fl. Ind. 2: 16 (1832). Type: Bangladesh, Chittagong, Wallich 1661a (lectotype K-W [as 1661.1], designated here; isotypes BM, BR, E, G, K, P). Echites reticulata Wall., Num. List 1662 (1829) pro parte nom. nud. [1662.2 in K - W].

EchitesacuminataRoxb., Fl. Ind. 2: 15 (1832) non Ruiz & Pav. (1799). Aganosmaacuminata G. Don, Gen. Syst. 4: 77 (1837); Wight, Ic. P1. 2: t. 424 (1843); Miq., Fl. Ind. Bat. 2: 447 (1857); Merr., Fl. Manila 374 (1912); Merr., Enum. Philipp. Fl. P1. 3: 334 (1923); Tsiang, Sunyatsenia 2: 156 (1934); Tsiang & Li, Fl. Rei. Pop. Sin. 63: 182 (1977); Guangdong Coll. Agric. For. (P. T. Li), Fl. Hainanica 3 (1974) 240; Ly, Feddes Repert. 97: 658 (1986). Ichnocarpus acuminatus Fern.-Vill., Novis App. 131 (1880).

Amphineurionacuminata (Roxb.) Pichon, Bull. Soc. Bot. France 95: 215 (1948). EchitesapoxysVoigt, Hort. Sub. Calc. 522 (1845) nom. illeg. Type: Illustration 2461 of Echitesacuminatain the Roxburgh collection in Kew library. EchitesreticulataBojer, Hort. Maurit. 211 (1837) non Roth in Roem. & Schult. (1819). A specimen collected by Bojerfrom a cultivated plant in Mauritius is in G - DC AganosmamacrocarpaA. DC., Prod. 8: 434 (1844); Miq., Fl. Ind. Bat. 2: 447 (1857). Ichnocarpus macrocarpus Fern.-Vill., Novis App. 131 (1880).

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FIG.4. Aganosma gracilis A Habit; B Flower; C Dissected flower. Drawn from Cooper1077 by Holly Nixon.

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471

MAP2. Distribution of Aganosma gracilis (0)

and Aganosma schlechteriana (0).

Echites macocarpa Wall., Numer.

List 1662 (1829)

nom. nud. Type: Wallich 1662

(lectotype G - DC, designated here; isotypes K, K - W [as 1662.1]) AganosmavelutinaA. DC., Prod. 8: 434 (1844); Miq., Fl. Ind. Bat. 2: 447 (1857); Merr., Enum. Philipp. Fl. Pl. 3: 334 (1923); Backer & Bakh. f., Fl. Java 2: 238 (1965). IchnocarpusvelutinusFern.-Vill., Novis App. 131 (1880). Amphineurionvelutinum(A. DC.) Pichon, Bull. Soc. Bot. France 95: 215 (1948). Type: Philippines, s.l., Cuming 1803 (lectotype G, designated here; isotypes E, G, K, L, MO, P, TCD, UPS, W). AganosmaeulobaMiq., Fl. Ind. Bat. 2: 447 (1857); Boerl., Handl. Fl. Ned. Ind 2: 400 (1899). Type: Indonesia, Java, Horsfields.n. (lectotype K, designated here; isotypes K, L [fragment], U). EchitesrepensBlanco, Fl. Filip. ed. 1, 109 (1837).

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MAP 3. Distribution of Aganosma marginata (0).

Echites procumbens Blanco, Fl. Filip. ed. 2, 78 (1845). Holarrhena procumbens (Blanco) Merr., Govt. Lab. Publ. (Philip.) 27: 59 (1905). Type: Not found. Neotype: Philippines, s.l., Merrill Species Blancoanae No. 372 (neotype K; isoneotypes BM, L, MO, P, US, W). Stems densely lenticellate, glabrous to short puberulent. Leaves: petiole 3 - 11 mm long; blade elliptic, oblong or narrow ovate, apex acuminate, base rounded to cuneate, 2.2 - 14 x 0.7 - 4.7 cm, 1.3 - 7 x as long as wide, 8 - 18 pairs of lateral before margin forming a strong nerves, prominent abaxially, anastomosing to then more densely abaxially especially nerve, intramarginal glabrous puberulent, on veins. Inflorescencelax, terminal and axillary, glabrous to sparsely puberulent, 420 cm long; pedicels 0.6 - 9.5 mm long. Sepals oblong to lanceolate, apex acute or acuminate, 2 - 8 x 0.8 - 2.1 mm, 1.7 - 5.3 x as long as wide, shorter than to as long as the corolla tube, puberulent to glabrous, with a continuous row of colleters on the

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473

inside. Corollawhite; tube 4.4 - 9.9 mm long; lobes linear, 5.5 - 21.9 x 1.2 - 3.3 mm, 3 - 9.5 x as long as wide, 1 - 2.8 x as long as tube; glabrous or minutely puberulent outside, short puberulent inside and densely puberulent just beneath stamen insertion. Stamensinserted at 2.2 - 4 mm from corolla base, 0.3 - 0.5 of tube length; anthers 2.5 - 3.2 x 0.4 - 0.7 mm, 3.7 - 7 x as long as wide. Disk of 5 wide rounded lobes to annular, 0.2 - 0.6 mm long, 0.3 - 1 x as long as ovary. Ovary 0.4 - 1.2 mm

long, minutely puberulent to glabrous; style 1.5 - 3.1 mm long; pistil head 0.8 - 2 mm long. Fruit long linear, lenticellate, glabrous to sparsely puberulent, 16.5 - 74 x 0.4 - 1 cm. Seeds 4.7 - 13.6 x 0.6 - 2.3 mm; coma 2 - 5.1 cm long. Fig. 5. Map 3.

DISTRIBUTION.India, Bangladesh, China, Burma, Thailand, Laos, Cambodia, Vietnam, Malaysia, Singapore, Philippines, Indonesia. HABITAT.Wide variety of habitats especially in drier evergreen and deciduous forest and as a scrambler in scrubland, to 850 m. Selection of the approx. 340 specimens studied: BANGLADESH.Chittagong, Wallich 1661 B (L, M, MEL); Sylhet, Wallich 1661a (BM, BR, E, G, K, P - Type of A. marginata). BURMA.Irrawaddy: Bassein, Rodger21546 (MEL). Karen State: Joukyeghat, Kurz 562 (M). Mandalay: Kyankpadamy, Papa Hill, Kyi 660 (A). Mon: Moulmein, Parish 191 (K). N Shan States: Lashio, Lace 5844 (E, K). Pegu: Prome, McClellands.n. (K, P). Rangoon: McClellands.n. (K). Tenasserim: Tavoy, Parish 158 (MEL). CAMBODIA.Siemreap: Angkor, Lecomte& Finet 1789 (K, P). Kampong Chhnang: Mueller 462 (P). Kampong Spoe: Pum Ho Tet, Poilane 17730 (P). Stung Treng: between Anlong Veng and Don Sa, Poilane 14969 (P). CHINA. Hainan: Yaichow, Chun & Tso44535 (A, B, K, P, US). INDIA. Andamans: Barren Isl., Kamphoevener 696 (C). Sikkim: Darjeeling, Cowan

s.n. (E).

INDONESIA.Flores: Wai Nuba, Malantarak, Afriastini 1585 (BO, L). Java: s.l., Horsfields.n. (K, L, U - type of A. euloba).Java Barat: Bogor, Nyman 50 (UPS). Java Timur: Kangean, Saboenten, Backer29928 (K, L). Sulawesi Selatan: Selayar, Teysmann 13872 (BO, K, L). Sumatera Aceh: Gunung Leuser National Reserve, Gunung Guhra, 15886 (BO, L, US, Z). Sumbawa Pernek, opposite Ketambe, de Wilde& de Wilde-Duyfjes Olat Seli, 12 km S of Sumbawa Besar, Kuswata224 (A, BO, K, L, SING). LAOS. Kham Mouan: Poilane 28186 (P). Louang Prabang: Tixiers.n. (P). Saravan: Poilane 15489 (P). Savannakhet: Poilane 28020 (P). Viang Chan: Barikhane, Kerr 21304 (BM, K, L, P). Xaigna Bouri: Ban Nale, Vidal4159 (P). MALAYSIA. Kedah: Pulau Langkawi Symington34720 (KEP). Melaka: Alor Gajah, 115 (SING, Z). Pahang: Taman Negara, ChengFRI 23320 (K, KEP,L). Penang, Berry Curtis664 (K, SING). Perak: Larut, King's Collector7321 (L, P). Sabah: Kota Belud, Melegrito3311 (K, L). Terengganu: Kuala Terengganu, Holttum15230 (SING). PHILIPPINES.Luzon: Albay, Cuming 1121 (BM, G, K, L, M, MEL, MO, P, TCD, UPS, W); Bataan, Mabesa & Sulit 26739 (A, BO, BRI); Bulacan, Mt Angat, Vidal459 (A, K, L); Cagayan, Vicinity of Penablanca, Adduru 97 (A, K, P, US); Central, San Francisco del Monte, Loher3993 (K, M, US); Isabela, Santiago, Clemens19000 (BO); Laguna, Canicosa 427 (A); Laguna, Calauan, Vidal M 1297 (AAU); Manila, San Mateo, Vidal 3286 (K); Nueva Vizcaya, near Dupax, McGregor 11224 (US);

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FIG. 5. Aganosma marginata A Habit; B Flower; C Dissected flower; D Fruit; E Seed. Flowering portion drawn from Poilane 14969, fruiting portion from Tixier s.n.

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475

Pangasinan, Umingan, Otanes 17986 (A, US); Rizal, Novaliches, Steiner 1022 (L, LAE); Zambales, Acoje Mine concession, Santa Cruz, Ridsdale & Reynoso1430 (A, BO, K, L). Masbate: Ticao, Clark1082 (US). Mindanao: Davao Del Sur, Todaya, Mt Apo, Elmer10998 (A, BM, BO, E, G, K, L, MO, P, US, WRSL, Z). Mindoro: Oriental, Puerto Galera, Santos 5260 (L, US). Mountain Province: Bontoc, Vanoverbergh 3863 (P). Negros: Cuming 1803 (E, G, K, L, MO, P, TCD, UPS, W - type of A. velutina). Palawan: Lipuun, Quezon, mouth of Bato Puti cave, Cordero& Espiritu 91493 (K, L); Puerto Princesa Mt Pulgar, Elmer12910 (A, BM, BO, E, G, K, L, MO, P, U, US, W, WRSL, Z). SINGAPORE. Murton 74 (K). THAILAND. Bangkok: Kerr 10674 (BM, K). Buri Ram: Phengkhlai et al. 3433 (BKF).

Chaiyaphum: Dat Don, Larsenet al. 31779 (AAU, BKF,K, L, P). Chanthaburi: Nong Ban, Larsen et al. 32243 (AAU, B, K, L). Chiang Mai: Bung Phrao, Chanthamuk130 (L, PSU). Chiang Mai: Doi Sutep, Garrett1199 (A, ABD, B, E, K, L, P, TCD). Chiang Rai: Muang Fang, Hosseus 609a (M). Chon Buri: Si Racha, Collins 260 (K). Chumphon: Kapur, Khantchai 1102 (K). Kanchanaburi: Wang Yai, Sai Yok, Charoenmayu409 (A). Krabi: Khao Satuw, Kerr12434 (ABD, BM, K). Lamphun: Me Tar, Winit240 (BM, E, K). Loei: Nam Nao National Park, Murata et al. T- 51766 (TI). Mae Hong Son: Khun Yuam, Nimanong & Phumsomsaeng 1795 (A, BKF, PSU). Nakhon Ratchasima: Sorensenet al. 2502 (C, PSU, Z). Phetchabun: Muang, Kerr5727 (ABD, BM). Phetchaburi: Poilane s.n. (A). Phrae: Hi Si, Vanpruk272 (K). Prachin Buri: Kabin District, Phengkhlai6827 (BKF, C, K, PSU). Prachuap Khiri Khan: Sam Roy Yot, Larsen & Larsen 33605 (C). Ranong: Khao Bangben, Kerr 16651 (AAU, ABD, BM, K). Ratchaburi: Lakshnakara405 (AAU, ABD, K). Rayong: Ko Samet, van Beusekom& Santisuk3249 (AAU, BKF,C, E, L, P). Saraburi: Sahm Lahn, Maxwell74276 (L). Satun: Adang Isl., Laem Sone area, Terutao Nat. Park, Maxwell87-325 (A, L, PSU). Si Sa Ket: Kantrarom, Suvanakoses 1578 (K). Songkhla: Khao Tao, Kerr 12686 (AAU, K). Trang: Chong, Boongird 69 (P). Trat: Ko Chang, Khlong Son, Sorensenet al. 7163 (C, Z). Ubon Ratchathani: Warin, Lakshnakara869 (ABD, BM, E, K, TCD). Udon Thani: Non Sang, Phu Khao, Bunchai 1549 (BKF,C, E, L, P). VIETNAM.Bien Hoa: Dong Nai near Tong Ben, Pierre1826 (A, BM, BO, C, K, L, MO, SING, US). Thuan Hai: Ca-Na, Poilane 5601 (A, BISH, P). Lam Dong: Da Lat, Squires819 (A, BISH, BM, BO, BR, G, K, M, MO, P, S, SING, US). Khanh Hoa: Nha Trang, Poilane 4511 (A, B, P, US). Phuoc Hoi: Suoi Da Tay Ninh, Mueller56 (P). Ho Chi Minh: Thorel867 (B, BO, K, P). Tay Ninh: Cai Cong, Pierre2026 (B, P). Trang Bom: Phung van Dieu 114 (P).

Hooker (1882) tentatively suggested that EchitescristataRoth in Roem. & Schult., cristata(Roth) G. Don, a new combination for Syst.Veg. 135 (1819) and Chonemorpha the same species, is a synonym of this species. If it were it would take priority over Echites marginata Roxb. (1832). I have found no Heyne specimen, on which Roth's descriptions are based, which could possibly be the type of this species. Although Roth's description of E. cristatafits A. marginatawell in a number of ways it would be premature and undesirable to displace the well established latter name for this common species without a definite type specimen for E. cristata.If it were established that these two species were synonymous A. marginatashould be conserved.

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VOL.51(3) KEWBULLETIN

Inflorescences which appear to be axillary are actually terminal with the continued growth of the vegetative shoot being formed from one of the axillary buds. 6. Aganosma schlechteriana Lev., Repert. Spec. Nov. Regni Veg. 9: 325 (1911); Hand.-Mazz., Symb. Sinicae 7: 991 (1936); Tsiang, Sunyatsenia 4: 31 (1939); Tsiang, Bull. Fan Mem. Inst. Biol. 9: 24 (1939); Guangdong Coll. Agric. For. (P.T. Li), Fl. Hainanica 3 (1974) 240; Tsiang & P. T. Li, Fl. Rei. Pop. Sin. 63: 185 (1977);J. Ying & P. T. Li, Fl. Yunnanica 3: 537 (1983); P. T. Li, J. S. China Agric. Coll. 11 (1990) 30. Type: Esquirol915 (lectotype E, designated here; isotype A [fragment & photo]). Syntype: China, Guizhou, Lao-Ten, Esquirol100 (E, A [fragment]) navaillei Lev., Fl. Kouy-Tcheou 32 (1914). Trachelospermum navaillei (Lev.) Tsiang, Sunyatsenia 4: 40 (1939); Tsiang & P. T. Li, Fl. Rei. Aganosma Sin. 63: 187 Pop. (1977);J. Ying & P. T. Li, Fl. Yunnanica 3: 539 (1983); P. T. Li,J. S. China Agric. Coll. 11 (1990) 31. Type: China, Guizhou, Lo Fou, Esquirol3653 (holotype E; isotype A [fragment & photo]). Aganonerionpoylmorphumauct. non Pierre ex Spire: Merr., Supp. List Hainan P1. 6:

330 (1930).

Aganosma odoraTsiang, Sunyatsenia 2: 154 (1934). Type: China, Hainan, Yaichow, How 70756 (holotype SYS[probably now in IBSC], n.v.; isotypes A, B, K, P, US). Aganosma cymosaauct non (Roxb.) G. Don: Tsiang, Sunyatsenia 2: 154 (1934); Rehder, J. Arnold Arbor. 15: 315 (1934); Rehder, J. Arnold Arbor. 18: 237 (1937). Aganosma radiata Merr., Lingnan Sci. J. 14: 51 (1935). Type: China, Hainan, Naam Shan Leng, Ngai District, Lau 309A (lectotype A, designated here; isotypes B, BM, E, G, K, P, US). AganosmamontanaKerr, Bull. Misc. Inform., Kew 1937: 93 (1937); Kerr, Fl. Siam. En. 2: 470 (1939). Type: Thailand, Nakhon Ratchasima, Khao Kuap, Put 2868 (lectotype K, designated here; isotypes A, ABD, BK, BM, E, L, P, TCD). var. brevilobaTsiang, Sunyatsenia 4: 34 (1939); Tsiang & P. T. Aganosmaschlechteriana Li, Fl. Rei. Pop. Sin. 63: 187 (1977);J. Ying & P. T. Li, Fl. Yunnanica 3: 539 (1983). Type: China, Yunnan, Szemao, Henry 11862 (holotype A; isotypes E, K, MO, NY). var. leptanthaTsiang, Sunyatsenia 4: 34 (1939); Tsiang & P. T. Aganosmaschlechteriana Li, Fl. Rei. Pop. Sin. 63: 187 (1977);J. Ying & P. T. Li, Fl. Yunnanica 3: 539 (1983). Type: China, Yunnan, Szemao, Henry 12047 (holotype A; isotypes K, MO, NY). Stems lenticellate or not; glabrous, tomentose or strigose. Leaves:petiole 4.3 - 17 mm long; blade elliptic or, rarely, obovate, apex acuminate to abruptly acuminate, base rounded

to acute, 3.1 - 13.5 x 1 - 6.9 cm, 1.5 - 3.6 x as long as wide, 4 - 10

pairs of lateral nerves, glabrous, sparsely strigose on petiole and midrib, or tomentose. Inflorescencea congested to lax terminal panicle, strigose or tomentose, 2.2 - 13 cm long; pedicels

1.8 - 22 mm long. Sepals narrow lanceolate,

6.3 - 15.2 x

1.5 - 5 mm, 1.8 - 7 x as long as wide, longer than corolla tube, tomentose, slender colleters in corners. Corollawhite; tube 4.8 - 12.7 mm long; lobes obovate, falcate, apex flattened or rounded, sometimes with one corner acute, 4.1 - 16 x 2.5 - 8.6 mm, 1 - 2.9 x as long as wide, 0.6 - 1.6(- 2.2) x as long as tube; short tomentose outside, densely pubescent in throat and in 5 rows inside. Stamensinserted at 1.2 3.9 mm from corolla base, 0.1 - 0.4 of tube length; anthers 3.8 - 6.3 x 0.8 - 1.3 mm,

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A REVISION OF AGANOSMA (APOCYNACEAE)

3.8 - 7 x as long as wide. Disk narrower at top, 5-dentate, 0.9 - 2.3 mm long, 1.1 - 2.9

x as long as ovary. Ovary0.6 - 1.6 mm long, glabrous to densely pubescent; style 1 2.8 mm long; pistil head 1.6 - 3.8 mm long. Fruit terete, glabrous to tomentose, 10.1 - 38 x 0.4 - 0.9 cm. Seeds 15.5 - 23 x 2.5 - 5.6 mm; coma 2.3 - 5.5 cm long. Map 2. DISTRIBUTION. India (Nagaland), Burma, China, Thailand, Vietnam. HABITAT. In mixed forest or thickets, 350 - 2800 m.

Selection of the 47 specimens studied: BURMA. Maymyo Plateau, Aui Sakau, Lace s.n. (E) CHINA. Guangxi: Napo County, Napo town, Qun w 5 (US); Tian'e Xian, Wang 43292 (A, MO); Hin Yeng, Ching 6742 (A); Na I, Ling Yun Hsien, Steward& Cheo 629 (A, BM, BO, G, S, W). Guizhou: Lao-Ten, Esquirol 100 (A, E - syntype of A. schlechteriana); Lo Fou, Esquirol 3653 (E, A (photo & scrap) type of Trachelospermum navaillei); Near Zhenning, Handel-Mazzetti 10351 (A, C). Hainan:

Yaichow, How 70756 (A, B, K, P, US - type of A. odora); Naam Shan Leng, Ngai District, Lau 309 A (A, B, BM, E, G, K, P, US - type of A. radiata); Loktung, Lau 26772 (A); Lin Fa Shan, Taam Chau-Lam Ko Dists., Tsang 307 (A, B, BM, E, G, K, MO, P, US, W). Yunnan: Yangbi Xian, Qingqiaohe stream, Bartholomewet al. 55 (A, E, US); To Kou region de Kioa Kia, Ducloux s.n. (P); Si-chour-hsien: Ping-chai, Feng 12549 (A); Schweli-Balwin divide, Forrest15926 (E, K); Mengtze, Henry 10806 (A, E, K, MO, US); Szemao,

Henry 12047

(A, K, MO -

type of A. schlechteriana var.

leptantha); Pien Oua, Ten 157 (E); Yung-jen Hsien, Tsai 52890 (A); Fo-Hai, Wang 73547 (A); Nan-Chiao, Wang75168 (A); Shunning, Wenkuankuai, Yu 16289 (A, E); Mienning, Mayetui, Yu 17700 (A, E). INDIA. Nagaland: Naga Hills, Khizobama, Bar 5096 (K) THAILAND.Nakhon Ratchasima: Khao Kuap, Put 2868 (A, ABD, BM, E, K, L, P,

TCD -

type of A. montana).

VIETNAM. Gia

Lai-Kon Tum: near Tu Mrang, Poilane 32284 (P, TCD).

7. Aganosma siamensis Craib,Bull. Misc. Inform., Kew 1915: 433 (1915); Pitard, Fl. Gen. Indo-Chine 3: 1225 (1933); Kerr, Fl. Siam. En. 2: 470 (1939); Tsiang, Sunyatsenia 4: 42 (1939); P. T. Li, J. S. China Agric. Coll. 11 (1990) 30. Type: Thailand, Chiang Mai, Doi Sutep, Kerr1797 (K lectotype, designated here; A, BM, E, K, P isotypes). Aganosma gracilis auct. non Hook. f.: Tsiang, Sunyatsenia 4: 37 (1939) pro parte; Tsiang, Bull. Fan Mem. Inst. Biol. 9: 23 (1939). AganosmagrandifloraMerr., J. Arnold Arbor. 23: 190 (1942); Ly, Feddes Repert. 97: 658 (1986). Type: Vietnam, Langsen, Thanh Moi, Petelot 2447 (lectotype A, designated here; isotypes MO, P, US). Aganosma kwangsiensisTsiang, Sunyatsenia 4: 37 (1939); Tsiang & P. T. Li, Fl. Rei. Pop. Sin. 63: 183 (1977); J. Ying & P. T. Li, Fl. Yunnanica 3: 537 (1983). Type: China, Guangxi, Tai-ming Shan, KwangsiMuseum415 (holotype IBSC). Aganosma kwangsiensis var. longilobata Y. Wan ex C. Z. Gao, Guihaia 10: 180 (1990).

Type: Wan87004 (GXMI holotype, n.v.). I have not seen the type specimen but the characters used to differentiate it fall within the range of variation of A. siamensis.

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Stems lenticellate, glabrous to sparselystrigose. Leaves:petiole 5 - 16 mm long; blade elliptic, oblong or obovate, apex acuminate, base cuneate to rounded, 3.9 - 14 x 1.6 - 7 cm, 1.7 - 3.4 x as long as wide, 4 - 10 pairs of lateral nerves, glabrous or with few hairs on petiole and midrib. Inflorescence lax and delicate, strigose and tomentose, 4.8 - 11.7 cm long; pedicels 3.5 - 19 mm long. Sepalslanceolate, 9.5 - 21 x 1.3 - 4 mm, 4.1 - 9.5 x as long as wide, longer than corolla tube, pubescent, few colleters in corners. Corolla white; tube 10 - 14 mm long; lobes narrow elliptic, apex acuminate or acute, 11 - 35 x 2.5 - 13 mm, 1.6 - 6 x as long as wide, 1 - 2.8 x as long as tube; puberulent outside, pubescent in throat and in 5 rows inside. Stamensinserted at 1.1 - 2.7 mm from corolla base, 0.1 - 0.2 of tube length; anthers 4.7 - 6.1 x 0.9 - 1.2 mm, 4.5 - 6.1 x as long as

wide. Disk narrower at top, 5-dentate, 1.4 - 3 mm long, 1.3 - 2.5 x as long as ovary. Ovary0.6 - 1.4 mm long, pubescent; style 1 - 1.8 mm long; pistil head 1.9 - 3 mm long. Fruitpuberulent, 14.2 - 16 cm long, 6 - 8 mm wide. Seedsnot seen. Map 4. DISTRIBUTION. China, Thailand, Vietnam.

In evergreen or deciduous forest, 500 - 1300 m. Guangxi: Tai-ming Shan, Kwangsi Museum 415 (IBSC-type of A. kwangsiense).Yunnan: Fabo, Banhong District, Cangyan County, Li 11666 (K); NanChiao, Wang76864 (A, UPS). THAILAND. Chiang Mai: Doi Chiang Dao, Khantchai342 (BKF,K), Phloenchit1108 (BKF,K), Khantchai406 (BKF,K); Chiang Dao District, Ban Yang Pong, Banziger675 (K); Doi Chiang Dao, SE foothills above Ban Yang Dong Luang, Maxwell 89-434 (AAU, BKF,L, MO), Maxwell90-464 (A, MO); Doi Sutep, Kerr1797 (A, BM, E, K, P - type of A. siamensis). Phrae: Nam Khrai, Huai Yuak, Smitinand & Cheke10798 (BKF,K, L, P). VIETNAM. Langsen: Thanh Moi, Petelot 2447 (A, MO, P, US type of A. 19687 Poilane Route Bien Nai: Hoa: Poilane 19687 20, (P). Lai (P). Dong grandiflora); Chau: Mu'o'ng Tong, Poilane25768 (P, TCD). Dong Nai: Route 20, Poilane 19687 (P). HABITAT. CHINA.

8. Aganosma wallichii G. Don, Gen. Syst. 4: 77 (1837); Kerr, Fl. Siam. En. 2: 470 (1939). AganosmacalycinaA. DC., Prod. 8: 432 (1844); Kurz, Forest Fl. Burma 2: 186 (1877); Hook. f., Fl. Brit. Ind. 3: 664 (1882); Watt, Products of India 1: 128 (1889); Brandis, Indian Trees 464 (1906); King & Gamble, J. Asiat. Soc. Bengal 74: 497 (1907); Koorders-Schumacher, Syst. Verz. 1 (1912) 181; Ridl., Fl. Mal. Pen. 2: 365 (1923); Tsiang, Sunyatsenia 4: 38 (1939). Echites calycinaWall., Cat. 1653 (1829) nom. nud. Type: Burma, Tavoy, Wallich1653 (lectotype K - W, designated here; isotypes BM, G - DC, K).

Echitescaryophyllataauct. non Roxb.: Blume, Bijdr. 1041 (1826) pro parte (excluding citation of Rheede drawing); Roxb., Fl. Ind. 2: 11 (1832). AganosmacaryophyllataG. Don, Gen. Syst. 4: 77 (1837). Aganosma blumeiA. DC., Prod. 8: 432 (1844) pro parte, Miq., Fl. Ind. Bat. 2: 446 (1857); Boerl, Handl. Fl. Ned. Ind. 2: 400 (1899); Tsiang, Sunyatsenia 4: 38 (1939); Backer & Bakh. f., Fl. Java 2: 237 (1965) nom. illeg. Type: Blume s.n. (lectotype U). (see nomenclatural discussion under A. dichotoma). Aganosmaroxburghiiauct. non G. Don: Wight, Ic. t. 440 (1841).

A REVISION OF AGANOSMA (APOCYNACEAE)

479

MAP4. Distribution of Aganosma siamensis (0) and Aganosma wallichii (0).

Stems lenticellate or not, stems strigose or tomentose, often becoming glabrous. Leaves: blade elliptic, oblong or weakly obovate, apex acuminate, base rounded or obtuse, rarely acute or weakly cordate, 4.5 - 15.5 x 1.7 - 8.5 cm, 1.2 - 3.5 x as long as wide, 5 - 13 pairs of lateral nerves, glabrous, sparsely strigose on petiole and abaxial midrib or tomentose. Inflorescencerobust and lax, strigose or tomentose, 4.6 - 19.5 cm long; pedicels 6 - 17 mm long. Sepals narrow ovate to lanceolate, 9 - 23 x 1.8 - 6.1 mm, 3.2 - 10 x as long as wide, longer than corolla tube, tomentose, colleters few in corners or frequently absent. Corollawhite or yellowish; tube 8.3 - 14 mm long; lobes obovate, falcate, apex flattened or rounded, sometimes with one corner acute or acuminate, 8.8 - 23 x 3.7 - 11.5 mm, 1.3 - 2.8 x as long as wide, 0.9 - 2.3 x as long as tube; tomentose or strigose outside, glabrous in throat, pubescent behind anthers inside. Stamens inserted 2.4 - 4.5 mm from corolla base, 0.2 - 0.4 of tube length; anthers 4.7 - 7.5 x 1 1.6 mm, 3.3 - 6.1 x as long as wide. Disk narrower at top, 5-dentate, 1.1 - 3 mm long, 1.1 - 3.8 x as long as ovary. Ovary 0.8 - 1.8 mm long, pubescent; style 1.9 - 3.5 mm long, sometimes pubescent; pistil head 2.2 - 3 mm long. Fruit unknown. Map 4.

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DISTRIBUTION. Burma, Thailand, Malaysia, Indonesia HABITAT. In evergreen forest, to 400 m.

(Sumatra, Java).

BURMA.Rangoon University, Parkinson14774 (K). Tenasserim: Tavoy, Paungdaw, Keenanet al. 1876 (E), Wallich1653 (BM, K - type of Aganosma wallichii); Mergui, Packmans.n. (K). INDONESIA. Java s.l. Blumes.n. (U), Lahaie s.n. (P), Lobb72 (K, TCD), Lobb1853 Unknown s.n. (K), (P), Unknowns.n. (L), Zippeliuss.n. (L). Java Barat: Depok, Hallier s.n. (BO, K, L), Burck s.n. (BO, L), Koorders41699 B (BO, L); NW of Bogor, G. Noentjoeng, Roempin, Saimoendt33 (BO, K, L, SING, U). Java Tengah: Pekalongan, Koorders27298 B (BO, K, L, P); Banyumas, Backer18886 (BO, K, L, P, SING), Koorders 22151 13(BO, L), Koorders41682 B (BO, L). Sumatera Aceh: Gunung Leuser Nature Reserve, Ketambe, Lau Alas valley, 35 km NW of Kutatjane, de Wilde& Wilde-Duyfjes 15684 (BO, K, KEP,L, MO, US). MALAYSIA. Johor: Bukit Durian, P. Pemanggil, Noor & Samsuri MN 54 (SING). Perak: Roekeyloe, King's collector10808 (BO, E); Scortechinis.n. (K). Selangor: Genting, Simpah, Strugnell 12891 (K, KEP, SING); Ulu Langat, Bukit Timbok, Sepandang Langat, Bubal KL 1595 (KEP); Ulu Gombak, Ng FRI 27356 (K, KEP); Kepong, Kochummen71969 (K, KEP,L, SING). THAILAND. Krabi: Khao Laeng Mai, Sangkhachand 1022 (BKF, C, K, L, P). Nakhon Si Thammarat: Lansagah, Karome Falls, Khao Luang National Park, Maxwell85-679 (A, AAU, BKF,E, L, MO, SINU). Pattani: Khao Kula Khiri, Kerr7747 (ABD, BM, E, K, L, P, TCD). Phatthalung: Khao Boo-Khao Yah National Park, Riang Tong Falls, Maxwell 86-525 (A, AAU, BKF, E, L). Prachuap Khiri Khan: Bang Saphan, Put 1454 (A, BM, K). Satun: Ko Terutao, Malacca Creek, Congdon 16 (A, AAU). Surat Thani: Khao Tao, Kerr 12969 (AAU, BM, K, P); Kanchanadit, Kerr 13055 (BM, K, SING); Ko Samui, Put 845 (BM, E, K, L, P, TCD); Na Sarn, Sanan 411 (BKF,K). Trang: Khao Chong, Maxwell75-888 (AAU, L). KNOWN TAXA INSUFFICIENTLY

Echites caryophyllatavar. glabrata Hassk., Cat. Hort. Bog. 123 (1844). Type: not designated. This variety could be a synonym of A. wallichii;it was described from material growing at the Bogor Botanic Garden in Java and A. wallichiiis the only species from Java which would fit the description. However, Hasskarl himself, in Flora 28: 300 (but incorrectly as 268) (1845), suggests it is a synonym of the Southern Indian and Sri Lankan A. cymosavar. elegans (but as A. elegans). This species was quite possibly cultivated in the garden at that time. Alternatively it could simply be the valid publication of the Wallich1651.2 specimen labelled in the Wallich herbarium as E. caryophyllatavar. glabra. This is an Indian plant synonymous with E. dichotoma. Physetobasis macrocarpa Hassk., Flora 40: 104 (1857). Holarrhena macrocarpa (Hassk.) Villar, Fl. Filip. 4: 130 (1880). De Kruif (1981) has this species as a synonym of Holarrhena pubescens (Buch.-Ham.) Wall. ex G. Don and, as a type specimen was not designated, gives a neotype for this taxon. The neotype, however, is given as a plant grown in the Pondicherry Botanic Garden in India despite the fact that Hasskarl specifically described a Javanese plant. Merrill

A REVISION OF AGANOSMA (APOCYNACEAE)

481

(1923) suggests that Holarrhena macrocarpaFern.-Vill., but not the Hasskarl basionym, is a synonym of Aganosma acuminata (= A. marginata). Certainly both the Physetobasis and the Holarrhena could be synonyms of Aganosma marginata

based on the descriptions but there is no type material on which to base this. What is very unlikely is that they are synonyms of Holarrhena pubescenswhich occurs in neither Java nor the Philippines. SPECIESEXCLUSAE Aganosma affinis G. Don, Gen. Syst. 4: 77 (1837). = Ichnocarpusfrutescens (L.) W. T.

Aiton Aganosma apoensis Elmer, Leafl. Philipp. Bot. 4: 1445 (1912). = Parsonsia curvisepala

K. Schum

Aganosma concanensis Hook., Ic. P1. ser. 2, 5: t. 841 (1851). = Parsonsia alboflavescens

(Dennst.) Mabb. Aganosma edithae Hance, Ann. Sci. Nat. Bot. Ser. 5(5): 227 (1866). = Cryptolepis elegans Wall. ex G. Don Aganosma laevigataJ. Graham, Cat. P1. Bombay 113 (1839). = Parsonsia alboflavescens

(Dennst.) Mabb. AganosmalaevisChamp. ex Benth., Hooker's J. Bot. Kew Gard. Misc. 4: 335 (1852). = Anodendron affine (Hook. & Arn.) Druce Aganosma petelotii Ly, Feddes Repert. 92: 628 (1981). = Parepigynum funingense Tsiang

& P. T. Li. This species is known only from one collection from Vietnam, Petelot 7203. The holotype is in Ho Chi Minh City and an isotype in the Missouri Botanical Garden. The corolla is infundibuliform and the calyx short and oblong. It is clearly not an Aganosma. I have synonymised it under Parepigynum funingenseTsiang & P. T. Li although I am not convinced that this genus is distinct from Beaumontia. Parepigynumis currently being revised by Yong-ping Yang in Wageningen and he may clear up this issue. Aganosma romburghii (Boerl.) Backer ex Tsiang, Sunyatsenia 4: 28 (1939) nom. illeg. = Urceolajavanica (Blume) Boerl. Aganosma roxburghii G. Don, Gen. Syst. 4: 77 (1837). = Kamettia caryophyllata (Roxb.) Nicolson & Suresh [as typified but not for the description].

ACKNOWLEDGEMENTS

I would like to thank the directors of the herbaria that provided material or hospitality during visits;John Parnell and Matthew Jebb for their comments on the manuscript; Holly Nixon for the illustrations and Marcella Campbell for help with the curation of material. This work was carried out with the financial support of the European Union Human Capital and Mobility Programme.

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KEWBULLETIN VOL. 51(3) REFERENCES

Bentham, G. & Hooker, J. D. (1876). Ichnocarpus. In: Genera Plantarum 2: 717. Reeve & Co., London. Blume, C. L. 1826. Bijdragen tot de flora van Nederlandsch IndiS. Lands Drukkerij, Batavia. Candolle, A. L. P. de (1844). Aganosmain Prodromus 8: 432 - 434. Paris. Don, G. (1837). Aganosma.In: A General History of the Dichlamydeous Plants vol. 4: 77. C. J. G. & F. Rivington, London. Holmgren, P. K., N. H. Holmgren & L. C. Barnett. 1990. Index Herbariorum. IAPT, New York. Hooker, J. D. (1882). AganosmaIn: Flora of British India 3: 663 - 665. Reeve & Co., London. Kruif, A. P. M. de (1981). A revision of Holarrhena R. Br. (Apocynaceae). Meded. Landbouwhoogeschool 81 - 2: 1 - 40 Leeuwenberg,

A. J. M. (1988).

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